Old sleeping Sicilian beauty: seed germination in the palaeoendemic Petagnaea gussonei (Spreng.) Rauschert (Saniculoideae,Apiaceae)

Petagnaea gussonei (Apiaceae) is a perennial herbaceous species endemic to northeast Sicily (Nebrodi Mountains). It is considered a remnant of the Sicilian Tertiary flora, and is endangered according to the Red List. There is no information in the literature about the germinability of its seeds, even though seed production is know to occur. The aim of this study was to obtain data to better understand seed germination of this species and its biological implications. Thus, several approaches were employed: vitality analyses, gibberellic acid supply, germination and soil microbial flora analyses via end‐point and qPCR. The results suggest that seed germination occurs after ca. 1.5 years at a rate of ca. 11%. The seeds can be classified as physiologically dormant, and probably require prolonged cold stratification for germination. Because seed germination is low, it is likely that agamic reproduction represents an important mean for its conservation and survival. These results have important implications for P. gussonei survival and should be considered in possible re‐introduction attempts aimed at restoring threatened populations.     

Reproductive success and clonal genetic structure of the rareArnica montana (Compositae) in The Netherlands

In a medium-sized population ofArnica montana, a threatened species in The Netherlands, the breeding system, reproductive success and genetic clonal structure were studied. Pollination experiments suggested thatA. montana is largely self-incompatible. Inbreeding depression was observed for seedling weight but not for fruit weight and germination rate. Although genetic variation is rather low in this population, the data suggest an outcrossing mating system. Analysis of the genotype of all mapped rosettes in a plot of 100 m² indicated that dense clusters often consist of identical genotypes, suggesting a clonal structure. Open clusters frequently contained several different genotypes. This may be caused by limited fruit dispersal, since seedlings were found mainly within or in the near surroundings of the clusters.     

Fruit fate,frugivory, and fruit characteristics: a study of the hawthorn,Crataegus monogyna (Rosaceae)

Summary The fate of fruits from a population of European hawthorn (Crataegus monogyna) in western Oregon, USA, was examined over a two-year period. Only one frugivore, the American robin (Turdus migratorius) foraged on the C. monogyna fruits, making this an unusually straightforward fruit-frugivore system. Dispersal efficiency was low, with an average 21% of seeds being dispersed (carried away from parent plants) each year; the most common fate of fruits was to simply fall. Robins dropped 20% of the fruits that they picked, and defecated/regurgitated 40% of the fruits (seeds) that they swallowed, beneath parent plants. One trait, gruit abundance, strongly affected the probability of bush visitation by robins, bushes with larger fruit displays being preferred. Both absolute dispersal success (number of seeds) and dispersal efficiency (proportion of seeds; success per propagule) were also found to be correlated most strongly with initial fruit abundance. Individual plant fecundity and fruit quality were found to vary little between years; as a result, dispersal efficiencies for individual plants were also annully consistent. Larger (older) plants produced more fruits and therefore had higher fitness. These results suggest that the optimal fruiting strategy for C. monogyna is therefore to get as big as possible as quickly as possible by delaying fruiting until later in life.     

REPRODUCTIVE ECOLOGY OF JEFFERSONIA DIPHYLLA (BERBERIDACEAE)

The reproductive ecology of Jeffersonia diphylla (L.) Pers. (Berberidaceae) was investigated by studying its breeding system, ovule production, seed set, seed dispersal by ants and seed predation by rodents. This species flowers early in the spring and is facultatively autogamous. In a typical year fruit and seed set is high (90%), however, freezing temperatures from late spring frosts in 1983 and 1985 resulted in low fruit set (7% and 20%, respectively), and reduced seed set in those flowers that produced fruit. No differences in seed set between selfed and outcrossed flowers were observed over a two-yr period (1983–84). Ovule number per capsule increased with plant size as measured by leaf number. Seed set and seed wt were unaffected by leaf number unless leaves were removed after flowering was initiated. Jeffersonia diphylla is myrmecochorous. Ants removed seeds faster when seeds were placed in areas where J. diphylla plants were absent, suggesting that dispersal within J. diphylla populations is ant limited. Moreover, fresh (1 day old) seeds were removed by ants faster than 3 day old seeds. Seed predation by rodents prior to dehiscence from capsules is heavy in large populations (85–90%), and apparently negligible in small populations. Predation of seeds that are released from capsules is heavy (approx. 66%), particularly at night. Overall, seed predators consume about 96% of the seed crop in well established populations, but probably much less in small young populations. Hence, seedling recruitment is likely to be higher in small populations, whereas ramet production from rhizomes is the primary mode of propagation in large ones. The evolution of autogamy, early flowering, and myrmecochory are discussed in light of the results of this study.     

BREEDING STRUCTURE OF A YUCCA FILAMENTOSA (AGAVACEAE) POPULATION

Yucca filamentosa and its species-specific pollinator, the yucca moth, Tegeticula yuccasella (Lepidoptera: Prodoxidae), form a relationship that is often cited as a classic example of a coevolved plant-pollinator mutualism. Observations of the moth's behavior have led to predictions that moth dispersal is relatively limited and that, as a consequence, the self-compatible Y. filamentosa should experience relatively high rates of self-fertilization. In contrast, analyses of its mating system indicated that Y. filamentosa was predominantly outcrossed. To better understand effective breeding patterns in Y. filamentosa populations, 10 polymorphic allozyme loci were investigated to analyze the breeding structure of a natural Y. filamentosa population. Analyses revealed that Y. filamentosa is predominantly outcrossed, has multiply sired fruits, and that each fruit was sired by a different set of pollen donors. The effective number of pollen donors per fruit ranged from 1.56 to 3.13, indicating that some correlated mating exists within fruits. Paternity analyses revealed that pollen moved from 6 m to 293 m (mean = 118 m) within the study population and that a minimum of 10% of the progeny were sired by pollen originating outside of the population. These results are discussed in the context of the yucca–yucca moth mutualism.     

Analysis of the genus Petagnaea Caruel (Apiaceae), using new molecular and literature data

In Southern Italy, an endemic monotypic genus belonging to family Apiaceae occurs: Petagnaea (P. gussonei), relict of Tertiary flora, belonging to subfamily Saniculoideae. At present, P. gussonei is an endangered species and is included in various lists of species deserving special protection. The genus belongs to scapose hemicryptophytes and shares a sciaphilous habitat (hygrophilous woodland). This study is aimed at doing a complete contribution about the evolutionary history of Petagnaea, using molecular markers as plastidial DNA (cpDNA), nuclear ribosomal DNA (rDNA) and data present in literature. We used nucleotide sequences from four regions of the chloroplast genome (rps16 intron, trnL^(UAA) intron, atpB-rbcL intergenic spacer, and partial matK gene) to investigate possible haplotypes in Petagnaea populations. To have an idea of the molecular relationships of all populations of P. gussonei, the internal transcribed spacer (ITS) sequences, already employed in recent studies, were obtained for 18 populations. These sequences in combination with other Saniculoideae ITS sequences available from GenBank have been used for a further phylogenetic analysis. The results agree with the current classification of Saniculoideae in placing P. gussonei in tribe Saniculeae, since P. gussonei is in basal position to Sanicula. According to intraspecific chloroplast DNA diversity, no different haplotypes were detected. In addition to molecular data, morphology, cytology, phytochemistry and conservation status have been considered in the discussion.     

Reproductive systems and low outbreeding barriers between Jacaranda cuspidifolia and J. mimosifolia (Jacarandeae,Bignoniaceae)

The genus Jacaranda shows notable karyotype stability and a prevailing self‐sterile breeding system with evidence of late‐acting self‐incompatibility in several species. However, some studies have indicated self‐compatibility in J. mimosifolia, a species cultivated worldwide in tropical and subtropical areas. Jacaranda cuspidifolia is a closely related species with natural distribution broadly overlapping with that of J. mimosifolia, and manual heterospecific pollination studies have indicated that these species are interfertile, but there is no report on the breeding system of the former. In this study, we used hand‐pollination experiments, pistil longevity, epi‐fluorescence and histological analysis of post‐pollination events to determine the breeding system of J. cuspidifolia. We also employed intra and interspecific crosses and seed germination tests to reevaluate the breeding system of J. mimosifolia and the inter‐fertility between the two species. Some fruits were initiated from self‐pollinated pistils in J. mimosifolia, but none of them reached maturity. On the other hand, complete absence of fruit development by self‐pollination was verified in J. cuspidifolia, while in situ pollen tube growth and histological analysis of post‐pollination events in selfed pistils revealed the characteristic ovule penetration, fertilization and endosperm initiation also observed in other bignoniaceous species with late‐acting self‐incompatibility. Low outbreeding barriers seem to operate between these species because reciprocal interspecific crosses and hybrid seed germination tests indicated they are bilaterally interfertile. However, fruit/seed production and seed germinability were significantly lower when pistils of J. cuspidifolia were pollinated with pollen of J. mimosifolia, compared with crosses in the opposite direction, which indicates a partial unilateral incompatibility. This result is discussed in the context of the possible occurrence of self‐compatibility in J. mimosifolia. The low level of incongruity operating between the two species also points to their recent evolutionary divergence.     

Implications of newly-formed seed-dispersal mutualisms between birds and introduced plants in northern California,USA

I examined the role of bird dispersal in invasiveness of three non-native plant species in California, USA: Triadica sebifera, Ligustrum lucidum, and Olea europaea. I selected these species because their invasiveness in California is uncertain, but a survey of ornithologists highlighted them as likely bird-dispersed. I quantified bird frugivory of these plants, compared them with a native species (Heteromeles arbutifolia), and explored the management implications of dispersal mutualisms for these and other incipient invasive plants. Fruit removal by birds was sufficient to permit spread for all study species. Seed dispersers (rather than seed predators) and pulse feeders (flocking species with potential for long distance dispersal) performed most fruit removal for the non-native species, a pattern indicative of an effective dispersal regime. The number of fruiting trees per stand was a significant predictor of bird visitation. Founding population size may thus be important in management of invasive, bird-dispersed plants. Disperser-defined niches were relatively narrow because a few disperser species performed the majority of fruit removal from study trees, but each fruit species was consumed by a variety of potential dispersers. This results in strong pairwise niche overlap between some plant species. Ordinated by bird use, study site-species combinations clustered more by geographic location than by plant species, emphasizing the opportunistic nature of bird foraging. None of the non-native focal plant species appears dispersal limited, and all have formed novel mutualisms in California. It is possible that these plants are now in lag phases preceding bird-mediated invasion. Consideration of bird dispersal when evaluating invasiveness is therefore an imperative.     

Diplochory in two Jatropha (Euphorbiaceae) species of the Monte Desert of Argentina

The potential explosive seed dispersal under controlled conditions and the dispersal by ants in natural populations are compared between two diplochoric species: Jatropha hieronymi Kuntze and J. excisa Griseb. The seeds of J. hieronymi are more than eightfold heavier than J. excisa seeds, and were explosively dispersed considerably further distances, reaching a maximum of almost 18 m. The differences in explosive dispersal distances between the two species seem to depend on both carpel wall thickness of the fruit and aerodynamic shape of the seed. Seed removal by ants was positively correlated with the presence of the elaiosome and was higher for J. excisa (83.6%) than for J. hieronymi (31.6%). Seed size was the major factor affecting the removal by ants, as only large bodied ants were able to transport the large seeds of J. hieronymi. The larger size and the higher oleic acid content of the elaiosomes of J. hieronymi seeds had no influence on the observed removal rates by ants. In contrast, ants transported the J. hieronymi seeds further distances than J. excisa seeds. Jatropha hieronymi distances achieved by both dispersal modes are in the range of the furthest distances described for a diplochorous species. Finally, the possible advantages of this dispersal mode in arid zones are discussed.     

Generative ontogeny in Tiquilia (Ehretiaceae: Boraginales) and phylogenetic implications

Tiquilia is very different from the other members of the Ehretiaceae (Boraginales) in many aspects of morphology and ecology. Because detailed knowledge about flower and fruit traits is necessary to reliably infer character evolution of and within Tiquilia, we investigated flower to fruit ontogeny in eight species of Tiquilia using light and electron microscopy. Tiquilia accumulated a number of autapomorphies such as the prostrate growth form, the lack of lateral and ventral bundles in the gynoecium, and the formation of nutlet‐like mericarpids as dispersal units instead of more or less succulent drupes. The internal architecture of the superior bicarpellate ovary resulted from the development of several secondary septa including apical, basal and false septa, as it has been reported also from other Boraginales. However, no character found in Tiquilia can be regarded as synapomorphic with any other taxon of the Ehretiaceae. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 112 , 520–534.     

Reproductive biology of Setaria magna Griseb. (Poaceae: Panicoideae: Paniceae)

The genus Setaria is economically important because many species are cultivated for grains or forage. Setaria magna is an American species, native to North America and introduced in South America. The morphological aspect of this species is similar to S. italica (foxtail millet), suggesting its potential value as a crop. The purpose of this work was to understand the breeding system (self-pollination vs. open pollination) of S. magna; additionally, the floral development was described. The results of the breeding system analysis indicated that S. magna is mainly autogamous and does not appear to have a self-incompatibility mechanism. The floral development observed was similar to that described for other Paniceae; in most of the spikelets only the upper anthecium developed fruit, but a small number of spikelets presented bisexual lower florets. These spikelets produced two caryopses per spikelet. Thus, S. magna can produced two types of dispersal units; a few whitish naked caryopses from the lower anthecium that fall without their lemma and palea at maturity; and a high number of brown caryopses protected by the lemma and palea from the upper anthecium. The presence of a normal embryo sac in the upper anthecium suggests that S. magna would present normal sexual reproduction, although we can not reject the formation of apomitic seeds in the lower anthecium.     

Temporal and spatial variation in dispersal in the greater flamingo (Phoenicopterus ruber roseus)

We studied movements of individually marked greater flamingos (Phoenicopterus ruber roseus) born in the Camargue, southern France, between their two most important breeding colonies in the western Mediterranean (Camargue and Fuente de Piedra, Spain) from 1986 to 1992. The two sites differ in the frequency with which they offer suitable conditions for breeding. Flamingos have bred each year in the Camargue since 1974, but in only 12 of the past 22 years at Fuente de Piedra. Higher colony fidelity is thus expected in the less variable environment (Camargue), but if dispersal occurs competition might be an important factor causing this dispersal. Following years during which breeding birds in the Camrgue were disturbed (1988 and 1990) a higher proportion of adults changed colonies between breeding attempts (= breeding dispersal, 12.4%), while only 0.4% of flamingos breeding in the Camargue dispersed in the other years. As expected, flamingos breeding at Fuente de Piedra showed a higher rate of breeding dispersal (8.14%). No differences were observed between males and females. The importance of breeding failure as a factor causing breeding dispersal in flamingos was also confirmed by the movements of individual birds. The proportion of young flamingos that moved from their natal colony to start breeding at Fuente de Piedra (= natal dispersal) was independent of sex and age, but increased when breeding access to the Camargue colony was more difficult. However, natal dispersal was also higher in 1988 and 1990 (40.5%) than in the remaining years (1.2%), as was breeding dispersal. We discuss possible ways in which the increased natal dispersal among inexperienced birds could be linked with the increased breeding dispersal of adults in the same year.     

The underdog invader: Breeding system and colony genetic structure of the dark rover ant (Brachymyrmex patagonicus Mayr)

Ants are among the most successful species at invading new environments. Their success undeniably comes from their various modes of reproduction and colony breeding structures, which influence their dispersal ability, reproductive potential, and foraging strategies. Almost all invasive ant species studied so far form supercolonies, a dense network of interconnected nests comprising numerous queens, without aggression toward non‐nestmates. This strategy results in invasive colonies that are able to grow extremely fast and large while avoiding intraspecific competition, allowing them to monopolize environmental resources and outcompete native species. Here, we developed and used 10 microsatellite markers to investigate the population structure and breeding system of the dark rover ant Brachymyrmex patagonicus Mayr in its introduced range. We determined whether this species exhibits a supercolonial structure by assessing whether different nests belonged to the same genetic colony. We inferred its dispersal ability by investigating isolation by distance and estimated the numbers of queens per colonies and mating per queen through parent‐offspring inferences. We found that most of the colonies of B. patagonicus were comprised of a single nest, headed by a single queen. Each nest was distinct from one another, without isolation by distance, which suggests strong dispersal ability through nuptial flights. These features are commonly observed in noninvasive and native ant species, but they are surprising for a successful invasive ant, as they strongly differ from other invasive ants. Overall, we discuss how this seemingly unfavorable strategy for an invasive ant might favor the invasive success of the dark rover ant in the United States.     

Disentangling the pollination system of the food-deceptive orchid Anacamptis longicornu (Orchidaceae): from breeding system to spatio-temporal variation in reproductive success

Abstract

The deceptive breeding system in the Orchidaceae family has been studied for decades to disentangle how pollination service can occur. Moreover, the interference of introduced herbivores on population development has been reported to affect the reproductive output in different plant species, including orchids. In this study, the breeding system of the food-deceptive species Anacamptis longicornu has been assessed. The spatio-temporal variation in both male and female reproductive fitness and its relationship with plant features were also analyzed. The possible herbivory effects were investigated by an exclusion experiment using two types of cages. The results confirm that this species is pollinator dependent and mostly allogamous but is also self-compatible. This species showed high values of cumulative inbreeding depression index for fruit set and seed viability parameters and was highly pollen-limited. Male fitness was always higher than female fitness between years and locations, and both parameters showed spatio-temporal variation. The use of cages excluded herbivores but, eventually, also excluded pollinators, so that both male and female reproductive success parameters decreased significantly in the complete exclusion treatment. The strong dependence of the species’ fitness on pollinator-mediated pollen transfer must be considered for the long-term conservation of A. longicornu.     

金沙江干热河谷滇榄仁开花物候与繁育系统

通过野外定点观测,并利用套袋实验、解剖实验以及花粉形态特征观察与活力检测、柱头可受性检测、花粉/胚珠比以及异交指数等,研究了金沙江干热河谷滇榄仁自然群体花部形态特征与开花进程、繁育系统特征以及传粉媒介与环境的适应性等内容。结果表明,滇榄仁始花期在每年4月上旬,群体花期持续时间为30d左右,开花同步性高,呈典型的"集中开花模式"。单花开放时,柱头先伸长和膨大,约4d后雄蕊伸出,具有雌雄异熟的特征,一定程度上避免了自交的发生。单花开放进程可划分为花蕾期、柱头伸长期、雄蕊始露期、盛开期、盛开后期、花谢期等6个时期。除了花蕾期外,整个开花期的花粉均具有活力。花粉活力与柱头可受性之间避开了最适授粉期,但也有部分重叠。滇榄仁可能同时具有风媒和虫媒传粉。繁育系统以异交为主,部分自交亲和,需要传粉者。滇榄仁花部特征、开花物候和繁育系统为其适应干热河谷恶劣气候环境提供了一定的生殖保障和进化潜力。良好的群落环境有助于促进滇榄仁异花传粉,产生更多有效的种子,从而促进林下自然更新。     

Pollination and Breeding System of two Co-Occurring Hirtella Species (Chrysobalanaceae) in Central Brazil

The phenology, major floral characteristics, breeding systems and fruiting success of two co-occurring species of Hirtella: H. glandulosa and H.gracilipes (Chrysobalanaceae) were studied in Central Brazil. The two species occur as trees in mesophyllous forests but H. glandulosa is frequent also in dense savanna areas. Both species flower at the end of the dry season (September) and have flowers with one-day longevity. In both species flowers produce nectar and are pollinated exclusively by butterflies. The number of visits recorded and pollination rate in each species were significantly different and indicate that pollinators prefer H. glandulosa flowers. Natural and controlled fruit sets were low in both species. The index of self-incompatibility (ISI) was 0.17 in H. glandulosa and 0.86 in H. gracilipes. Although ISI in H. glandulosa could denote an incompatibility system, the presence of fruits with aborted embryos at different degrees of development point to an inbreeding depression situation. Low pollen viability and fruit set in H. gracilipes suggested reproductive problems which may be linked to hybridization events.     

The fig wax scale Ceroplastes rusci (Linnaeus) (Homoptera: Coccidae) in south-east Vietnam: Pest status, life history and biocontrol trials with Eublemma amabilis Moore (Lepidoptera: Noctuidae)

The fig wax scale Ceroplastes rusci (Linnaeus) is a serious pest of fruit trees in many countries. In the present study we investigated the extent of C. rusci infestation and the range of host species, particularly in fruit orchards, in south‐east Vietnam. Captive populations of C. rusci were established to record life history parameters and to investigate the efficacy of an endemic predacious moth, Eublemma amabilis Moore, as a potential biocontrol agent. Heavy infestation of C. rusci (up to 100%) was recorded in soursop and other fruit orchards, and the culturally important Hoa Mai flower was also affected. Captive breeding trials found E. amabilis to be an efficient predator of C. rusci, but an unusual hyperactive trait in early instars of E. amabilis resulted in lower than expected survival rates. The implications of this trait in terms of the laboratory environment, augmentative release protocols and as a survival strategy are discussed.     

Differential diversifications of South American and Eastern Asian disjunct genera Bocconia and Macleaya (Papaveraceae)

Bocconia (10 species) and Macleaya (2 species) are two disjunct genera between South America and eastern Asia (EAS) in the Papaveraceae offering an opportunity to compare its biogeographic history with that of the well‐known disjunction between EAS and eastern North American (ENA). Our phylogenetic analyses of the chloroplast matK and rbcL gene sequences of Ranunculales including two species of Macleaya and six species of Bocconia supported the monophyly of Bocconia, Macleaya, and Chelidonioideae to which Bocconia and Macleaya belong. Nucleotide sequences of matK, rbcL, and nrDNA ITS supported the sister relationship of Bocconia and Macleaya. Biogeographic analyses of Chelidonioideae using S‐DIVA (statistical dispersal vicariance analysis) and DEC (dispersal extinction cladogenesis) methods inferred Eurasia as the most likely ancestral area of Bocconia and Macleaya and suggested no extinction events in either Bocconia or Macleaya. This agrees with the “Out‐of‐Asia” pattern of the EAS‐ENA disjunction. Molecular dating of Ranunculales with fossil‐based calibrations showed that Bocconia and Macleaya diverged in the late Eocene and early Oligocene, which is much earlier than most EAS‐ENA disjunct taxa. The disjunction may have formed via long distance dispersal or boreotropical connections via the North Atlantic and Bering land bridges. Both Bocconia and Macleaya diversified in the late mid‐Miocene, but Bocconia has apparently experienced a greater diversification probably aided by the evolution of the bird dispersal syndrome in fruit and seed after migration to South America. The greater diversification of Bocconia is also evidenced by the diverse leaf morphology and growth habit in response to colonization in various local habitats in South America.     

Breeding behaviour of Kunzea pomifera (Myrtaceae): self-incompatibility, intraspecific and interspecific cross-compatibility

To examine breeding system characteristics of the endemic Australian prostrate shrub Kunzea pomifera, artificial hybridisations were undertaken using thirteen different genotypes of K. pomifera, to elucidate: (1) self-incompatibility, (2) intraspecific cross-compatibility in the species and (3) interspecific cross-compatibility with each of K. ambigua and K. ericoides. K. pomifera exhibited very low self-compatibility, with the barrier to self-fertilisation being prevention of pollen-tube growth in the style or ovary. Following intraspecific pollination amongst a number of different genotypes of K. pomifera, 38.4% of pollinated flowers developed fruit; arrest of compatible pollen-tubes in the style, preventing fertilisation, contributes to the low fruit set in this species. Interspecific compatibility was examined between K. pomifera (pistillate parent) and K. ambigua (staminate parent) where seed set per pollinated flower (4.47) was not significantly different from intraspecific crosses (4.66). In crosses between K. pomifera (pistillate parent) and K. ericoides as staminate plant, 0.037% of pollinated flowers produced fruit, with 0.0075 seeds per pollinated flower. Reproductive barriers between these two species were evident in the style of K. pomifera, where the growing tips of the K. ericoides pollen-tubes swelled and ceased to grow.